The Steppe Elephant Mammuthus trogontherii (Pohlig) from the … · 2020. 2. 28. · The lower jaw...

ISSN 0031�0301, Paleontological Journal, 2015, Vol. 49, No. 3, pp. 304–325. © Pleiades Publishing, Ltd., 2015.Original Russian Text © A.V. Shpansky, S.K. Vasiliev, K.O. Pecherskaya, 2015, published in Paleontologicheskii Zhurnal, 2015, No. 3, pp. 81–102.

304

INTRODUCTION

Three skeletons of ancient elephants have beenrecorded in the West Siberian Plain. All of them areconfined to the basin or valley of the Irtysh River. Askeleton found on the Om River (right tributary of theIrtysh) at the village of Ust’�Tarka is the first record ofthe steppe elephant skeleton in Western Siberia. In1993, the second skeleton was found on the IrtyshRiver at the village of Chembakchino Hunty–MansiAutonomous Region (Kosintsev et al., 2004); and thethird skeleton was found in 2002 on the Irtysh River atthe town of Pyatiryzhsk of the Pavlodar Region(Shpansky et al., 2008a). In the skeleton from Pyat�iryzhsk, the skull is not preserved and, in the specimenfrom Chembakchino, the skull is only represented byseveral fragments. The skeleton from Ust’�Tarka has awell�preserved skull, which is important for the studyof morphology and evolution of Pleistocene elephantsin Western Siberia. This skeleton is described in thepresent study.

The skeleton was excavated by seasonal workers inthe summer season of 1989 in the river channel.Approximate coordinates of the record are 55°32′ N,75°41.5′ E (Fig. 1). Initially, as the skeleton was takenfrom a sand layer rich in water, it was probably com�plete. Subsequently, bones of the forelimbs (except forthree wrist bones) have been lost. Due to I.E. Greb�nev’s efforts, the preserved part of the skeleton wasbrought to the Institute of Archeology and Ethnogra�phy of the Siberian Branch of the Russian Academy ofScience, Novosibirsk (IAE) in 1990. At present, the

reconstructed and mounted skeleton of the steppe ele�phant is exhibited in the hall of IAE (Fig. 2).

TAPHONOMY AND GEOLOGY

Direct data on the burial conditions of the skeletonin question are absent, although it is possible to judgecertain burial conditions from the bone preservation.The bones are heavy, mineralized considerably; thesurface is impregnated with manganese hydroxide forup to 1–2 mm of depth. The surface of some bones wasinitially covered with a cemented sandy cover, whichstripped gradually off in the course of drying. It is plau�sible that the elephant cadaver was buried rather rap�idly under a fine�grain sand layer at the border of theriver channel. The skull preserved with tusks and lowerjaw suggests that the cadaver was not transported for along distance.

A.V. Shpansky investigated the presumable localityin 2002. The section, where the specimen was foundhad not been investigated, because it was completelyturf�covered. The nearest outcrop is located on the rightbank of the Om River, 2.5 km (along a straight line)downstream from the village of Ust’�Tarka. The dis�tance from the point of skeleton finding is several hun�dred meters. The following beds are recognized down�ward in the section: (1) soil (0.3 m thick); (2) yellowishbrown fine�grain, horizontally thin�layer, dense sand;in the upper part, sand passes into siltstone, with rarevertical fracturing; in the lower part, sand is medium�grained, encloses shells of the freshwater mollusk Ame�

The Steppe Elephant Mammuthus trogontherii (Pohlig) from the Irtysh Region Near Omsk

A. V. Shpanskya, S. K. Vasilievb, and K. O. Pecherskayaa

aTomsk State University, pr. Lenina 36, Tomsk, 634050 Russiae�mail: [email protected]; [email protected]

bInstitute of Archeology and Ethnography, Siberian Branch, Russian Academy of Science, pr. akademika Lavrent’eva 1, Novosibirsk, 630090 Russia

e�mail: [email protected] May 14, 2012

Abstract—A Pleistocene elephant skeleton was found in 1989 in the alluvial beds of the Tobolsk age on theOm River near the village of Ust’�Tarka (Ust’�Tarkskii District, Novosibirsk Region). The forelimbs andsome thoracic and caudal vertebrae are not preserved. Judging from the heavy wear of M3/m3, this animalwas about 50–60 years of individual age. The skeleton is about 3.5 m high at the withers. The geological ageof the beds enclosing the skeleton and its morphological features suggest that it should be assigned to Mam�muthus trogontherii.

Keywords: Steppe elephant, morphology, skeleton proportions, Middle Pleistocene, Western Siberia

DOI: 10.1134/S0031030115030107

PALEONTOLOGICAL JOURNAL Vol. 49 No. 3 2015

THE STEPPE ELEPHANT MAMMUTHUS TROGONTHERII (POHLIG) 305

soda asiatica (Marthens, 1874) and carbonate nodules(7.0 m thick); (3) brown with light blue spots, thin�interbedding clays, containing abundant carbonateconcretions (6.0 m of exposed thickness).

The lower part of the outcrop was closed by a land�slip and turf�covered, about 20 m thick. A similar tex�ture of Quaternary deposits is observed in high terracesthroughout the lower reaches of the Om River. It isprobable that the elephant skeleton comes from a ter�race base, where clays of Bed 3 are underlain by sandof the Tobolsk Horizon of the Middle Neopleistocene.

MATERIAL AND METHODS

Specimen IAE, no. 18, skeleton of male Mam�muthus trogontherii Pohlig, including a skull withtusks, lower jaw, seven cervical and 15 thoracic verte�brae (vertebrae 6–8 and 19 are lost), 29 ribs, sternum,last lumbar vertebra, sacrum, pelvis, both hind limbs,including almost all skeletal foot elements. Caudalvertebrae are not preserved.

The skull and lower jaw were measured followingthe method developed by Dubrovo (1960) and supple�mented by some measurements of vertebrae (Shpan�sky et al., 2008a) (Fig. 3) and long limb bones (Baigu�sheva and Garutt, 1987). The generation of function�ing molars is determined following Sher and Garutt(1985). The sex is determined based on the long diam�eter of tusks and pelvic structure, following Lister(1996b). In long limb bones of the steppe elephant,sexual dimorphism is poorly pronounced (Shpanskyet al., 2008b). The ranges of measurements of thescapulae and long limb bones of males and femalesoverlap considerably (Fig. 4). Only the indices of pel�vic bones (Table 10) calculated according to Lister(1996b) differ distinctly. Since the skeleton ismounted, it was impossible to measure or describe thebones of feet and some vertebrae. Some skull measure�ments are approximate, because its rostral part (pre�maxillae) was reconstructed.

Published and original data on nine skeletons ofM. trogontherii and also one skeleton and two skulls ofM. trogontherii chosaricus Dubrovo were used for com�parison (Table 1).

As the taxonomic position of the elephant fromUst’�Tarka was determined, we followed the point ofview traditional for Russian researchers concerningthe recognition of two genera among Euroasian mam�moth�like elephants, Archidiskodon Pohlig, 1885 andMammuthus Brookes, 1828 (Garutt, 1998; Garutt andTikhonov, 2001; Baygusheva and Titov, 2012). Theanalysis of morphological skull characters is based onpublished data on A. meridionalis, specimen SM,no. 14120 from the town of Georgievsk (Garutt andSafronov, 1965) and Italy: specimen IGF, 1054 fromUpper Valdarno (Palombo and Feretti, 2005); fromSauvignon (Reggiani and Sala, 1994). The material ofM. primigenius includes a skull (specimen PM TGU,

no. 1/54) and lower jaws (specimens PM TGU,nos. 1/34 and 1/56).

The following abbreviations are used in the presentstudy: (AMZ) Azov Reserve Museum, Azov, Russia;(IAE) Institute of Archeology and Ethnography of theSiberian Branch of the Russian Academy of Science,Novosibirsk, Russia; (IEI) Museum of Archeologyand Ethnography of the Institute of EthnographicStudies of the Ufa Scientific Center of the RussianAcademy of Science, Russia; (MP PGPI) Museum ofNature of the Pavlodar State Pedagogical Institute,Pavlodar, Kazakhstan; (PM OGU) PaleontologicalMuseum of Odessa State University, Ukraine;(PIN) Paleontological Museum of the Borissiak Pale�ontological Institute of the Russian Academy of Sci�ence, Moscow, Russia; (PM TGU) PaleontologicalMuseum of Tomsk State University, Tomsk, Russia;(TOKM) Tomsk Regional Museum, Tomsk, Russia;(KhM) “Museum of Nature and Man,” Khanty�Mansiisk, Khanty–Mansi Autonomous Region,Yugra, Russia; (IGF) Museo di Geologie e Paleonto�logia, Universita di Firenze, Italia; (SM) StavropolState Historical and Cultural and Nature–LandscapeReserve Museum, Stavropol, Russia. The symbol “C”preceding a measurement in the tables designates thatit is incomplete; The upper and lower molars of the lastgeneration are designated M3 and m3, respectively.

Ob River

Khanty–Mansiisk

Irtysh River

Chembakchino

Ust’�TarkaOm River

Omsk

R u s s i a

Kazakhstan

Novosibirsk

Pyatiryzhsk

Pavlodar

Localities of Mammuthus trogontherii Pohlig

Irtysh River

Fig. 1. Geographical position of localities of steppe ele�phant skeletons within the West Siberian Plain.

306


SHPANSKY et al.

Fig. 2. A skeleton of the steppe elephant Mammuthus trogontherii, specimen IAE, no. 18; Novosibirsk Region, Ust’�Tarka; MiddleNeopleistocene; mounted and exhibited in the hall of the Institute of Archeology and Ethnography of the Siberian Branch of theRussian Academy of Science (Novosibirsk).

1

4

5

6 7

8

9

10

10

Fig. 3. Scheme of measurements of the thoracic vertebrae of the steppe elephant (after Shpansky et al., 2008a): (1) greatest widthat transverse processes; (2, 4) cranial and caudal width of vertebral centrum; (3, 5) cranial and caudal height of vertebral centrum;(6, 7) cranial width and height of spinal canal; (8) neural spine length along anterior edge; (9) minimal width at articular rib fossaecranially anf caudally; (10) vertebral centrum length doesallt and ventrally.



DESCRIPTION

Skull (Pl. 10, figs. 1, 2; Table 2) without tusk alve�oli. The occiput apex and occipital crest is rounded intransverse outline. The occipital plane is convex andsmoothly anteriorly rounded in the upper one�third.The occipital condyles project significantly beyondthe occipital plane. The skull is high (for measure�ments and ratios, see Table 2). The forehead surface isslightly concave in the sagittal plane. The forehead israther wide. The zygomatic arch is positioned at anangle of 105° to the occipital plane in projection on thesagittal plane. The angle of inclination of the foreheadplane relative to the premaxillary surface is 150°. Skullwidth at ptojections is greater than zygomatic widthand greatest occipital width. The nasal process is trian�gular, with a sharpened lower edge, slightly projectsabove the level of the frontal surface. The nares aresemilunar, with the lateral edge extending stronglydownwards. The lower edge of the nares is slightlyshort of reaching the orbital midline. In Archidisk�odon, the nares are located in line with the upperorbital border (Baigusheva and Garutt, 1987). Theskull measurements of the elephant from Ust’�Tarkaare inferior to that of the Azov 1 skull and superior tothat of M. trogontherii chosaricus (Garutt, 1972;Shpansky, 2000) (Table 2).

The lower jaw (Pl. 10, figs. 3, 4; Table 3) has mas�sive horizontal rami strongly widened in the middlepart. The mental protuberance is long, flattened in the

sagittal plane, and pointed. The lower jaw of thefemale from the Azov 2 locality has a short mental pro�tuberance (Baigusheva, 2001). The anterior ends ofthe crista mentalis are well pronounced, ridgelike,with a relief surface. The left horizontal ramus has twolarge mental foramina, which are located under theanterior edge of the alveolus of m3. The right ramushas one larger mental foramen. The horizontal ramusmeasured along the anterior alveolar edge is muchdeeper than that measured along the posterior edge.The distance between the lateral edges of the articularheads is less than the width between the lateral surfacesof jaw corners. In M. trogontherii, the ratio of thesemeasurements is 89–90% (Table 3). The lower jaw ofthe male from Azov 1 shows an inverse relation. Baigu�sheva and Garutt (1987, table 2) provide a consider�ably overestimated measurement (in our opinion, by200 mm) of “the distance between the external edgesof jaw angles” (820 mm), which does not correspondto the value provided by these authors for the ratio ofthis measurement to the distance between the edges ofheads of the articular processes (101.2%).

The teeth belong to the last generation, M3/m3(Pl. 10, figs. 3, 5; Tables 4, 14). All plates are worn; thetalons are at the initial wear stage; several anteriorplates are already lost. The enamel plates are wide,anterior and posterior walls of the plates are moder�ately worn, positioned parallel to each other; thespaces between them are very small. The enamel is

35

30

25

20

15

10

5

4

3

2

1Scapula Humerus Ulna Radius Femur Tibia Pelvis

OdessaAzov 1Azov 2Ust’�TarkaPyatiryzhsk

ChembakchinoEderslebenWest RuntonOr’yaNovogeorgievsk

Fig. 4. Relationships of the extent of gracefulness of limb bones of steppe elephants (in the scapula, the ratio of the neck width tothe greatest bone length is used; in other bones, the ratio of the diaphyseal width to the greatest length; the data are marked usingthe left scale; for the pelvis, the index 3 : 5 (Table 10) is used, marked using the right scale).

308


SHPANSKY et al.

coarsely plicate; in the lower teeth, it is thicker than inthe upper teeth. In the crowns of M3, nine plates arepreserved. Right m3 retains seven plates and left m3has eight. The crown of the right tooth is somewhatmore raised above the alveolus; the left tooth is lowerby 0.5–1.0 cm. The upper teeth are located in accor�dance with them; the right tooth projects from thealveolus to a lesser extent than the left tooth, thus,compensating the difference in height of m3.

The alveolar parts of tusks about 1.5 m of length arepreserved. The tusks are circular in cross section,about 270 mm in diameter in the alveolar part and860 mm in circumference at the alveolar edge.

The total length of mounted cervical vertebrae is500 mm. The wing width of the atlas of the elephantfrom the Ust’�Tarka locality is less than that of ele�

phants from Azov and Pyatiryzhsk (Table 5). The neu�ral arch is horizontal, has a sinusoidal longitudinalconcavity on the caudal side. The dorsal arch has cres�tlike tubercles, which are particularly well developedin the lateral parts. The vertebra is relatively high, theratio of the vertebral height to greatest width at thewings is 59.3% (in the males from Azov 1 and Pyat�iryzhsk, it is 58.6 and 48.4%, respectively). In the mid�dle of the ventral arch, the vertebra has a well�devel�oped tubercle, the apex of which is displaced caudallyrelative to the transverse plane.

Epistropheus (or axis) (Table 6). The crest on theneural arch is divided by a deep sagittal notch and nar�rows cranially. The bases of the lateral parts of the cra�nial articular facets overhang transverse foramina. Thecaudal facet is slightly concave, widely oval. The spinal

Table 1. Skeletons of mammoth�like elephants involved in comparison

Locality (reference) Depository, no. Sex Geological age

Mammuthus trogontherii (Pohlig 1885)

Khadzhibeiskii estuary, Odessa, Ukraine (Yatsko, 1948)

PM OGU Male Early Neopleistocene

Novogeorgievsk, right bank of Dnieper River, Ukraine (Zakrevs’ka, 1935)

Not specified Female Not specified

Kagal’nitskii quarry, Azov, Rostov Region (Baigusheva and Garutt, 1987)

AMZ no. KP�21081 Male Not specified

Kagal’nitskii quarry, Azov, Rostov Region (Baigusheva and Titov, 2010)

AMZ no. KP�28689 Female Early stage of Tiraspol Assemblage, MQR6�5

Ust’�Tarka, Om River, Novosibirsk Region (original data)

IAE, no. 18 Male Tobolsk horizon, ca. 400 ka

Pyatiryzhsk, Irtysh River, Pavlodar Region, Kazakhstan (Shpansky et al., 2008a)

MP PGPI no. P2002.1149 Male Tobolsk horizon, ca. 400 ka

Chembakchino, Irtysh River, Khanty�Mansi Autonomous Region (Kosintsev et al., 2004)

KhM�10398 Male Second part of Early Neopleis�tocene, 550–600 ka

Gelsenkirchen, Westfalen, Germany (Siegfried, 1956)

Geology Museum, Münster Male Saale�Vistula Interglacial, begin�ning of Middle Neopleistocene, ca. 400 ka

Edersleben, Germany (Garutt and Nikolskaya, 1988)

Sangerhausen Female Beginning of Early Neopleistocene, ca. 700 ka

West Runton, Norfolk, Great Britain (Lister and Stuart, 2010)

Museum of Norfolk Life, no. 1992.36; 1996.181

Male ca. 700 ka

Mammuthus trogontherii chosaricus (Dubrovo 1966)

Starokudashevo, Or’ya River, Bashkiria (Garutt, 1972)

IEI, OF�909 Female Dnieper horizon, ca. 300 ka

Chernyi Yar, Volgograd Region (Dubrovo, 1966)

PIN, no. 4874 Male Dnieper horizon, ca. 300 ka

Town of Asino, Chulym River, Tomsk Re�gion (Shpansky, 2000)

TOKM, no. 10300/3 Male Samara horizon, ca. 300 ka

E x p l a n a t i o n o f P l a t e 1 0

Figs. 1–5. Mammuthus trogontherii (Pohlig 1885), specimen IAE, no. 18; Novosibirsk Region, Ust’�Tarka; the middle Neopleis�tocene, the Tobolsk horizon: (1) skull, anterior view; (2) skull, lateral view; (3) lower jaw, right lateral view; (4) lower jaw, dorsalview; (5) right upper М3, view of masticatory surface.



Plate 10

10 cm

10 cm

10 cm

5 cm

1

2

3

4

5

310


SHPANSKY et al.

Tabl

e 2.

Sku

ll m

easu

rem

ents

of M

amm

uthu

s tr

ogon

ther

ii (

Poh

lig,

188

5)

No.

Mea

sure

men

ts (

mm

) an

d in

dice

s (%

)

M. t

rogo

nthe

rii

M. t

rogo

nthe

rii c

hosa

ricu

s

Ust

’�Ta

rka,

spec

imen

IA

E,

no. 1

8

Azo

v 1,

spe

cim

en

AM

Z, K

P�2

1081

(Bai

gush

eva

and

Gar

utt,

198

7)

Gel

senk

irch

en

(Sie

gfri

ed, 1

956)

Or’

ya, O

F�9

09

(Gar

utt,

197

2)

Asi

no, T

OK

M,

no. 1

0300

/3(S

hpan

sky,

200

0)

Che

rnyi

Yar

,sp

ecim

en P

IN,

no. 4

874

(Dub

rovo

, 196

6)

1S

kull

leng

th: d

ista

nce

from

ant

erio

r or

bita

l bor

der

to o

ccip

ital

con

dyle

s in

clus

ive

840

695

780

600

775?

680

2P

arie

tal l

engt

h: fr

om a

pex

to a

nter

ior

edge

of p

rem

axill

ae14

50C

1910

1220

1190

1390

1390

3C

ondy

loba

sal l

engt

h: fr

om u

pper

(po

ster

ior)

edg

e of

occ

ipit

al c

ondy

les t

o lo

wer

(an

teri

or)

edge

of p

rem

axill

ae11

4014

4010

2010

6111

00

4F

oreh

ead

leng

th: f

rom

ape

x to

low

er e

dge

of n

asal

s (i

n pr

ojec

tion

on

mid

line)

670

5F

oreh

ead

leng

th a

long

wit

h na

res:

from

ape

x to

low

er e

dge

of n

ares

700

965

533

C58

058

057

0

6L

engt

h of

pre

max

illae

: fro

m lo

wer

edg

e of

nar

es to

low

er e

nd o

f pre

max

illae

850

962

750

812

870

7N

ares

(gr

eate

st w

idth

× g

reat

est h

eigh

t)63

0 ×

158

c700

504 ×

144

470 ×

160

460 ×

195

532 ×

155

8Z

ygom

atic

wid

th: g

reat

est s

kull

wid

th a

t zyg

omat

ic a

rche

s79

010

5070

078

5

9G

reat

est o

ccip

ital

wid

th83

510

5678

067

6C

760

805

10W

idth

of p

rem

axill

ae, u

pper

: at l

evel

of l

ower

edg

e of

infr

aorb

ital

fora

min

a (a

long

wit

hpr

oces

ses

of m

axill

ae)

495

640

420

430

350

456

11G

reat

est o

ccip

ital

wid

th a

t low

er e

dge:

bet

wee

n ex

tern

al e

dges

of t

usk

alve

oli a

t low

er r

os�

trum

end

__

_73

167

046

4~

560

530

12M

inim

um fo

rehe

ad w

idth

: min

imum

dis

tanc

e be

twee

n cr

ests

bor

deri

ng te

mpo

ral f

ossa

e43

064

0?36

031

031

0

13Tu

sk d

iam

eter

at a

lveo

lar

edge

(in

tern

al a

lveo

lar

diam

eter

)27

021

5 ×

174

160

(146

)(1

65)

191

14D

ista

nce

from

sku

ll ap

ex to

mas

tica

tory

mol

ar s

urfa

ce a

long

per

pend

icul

ar to

this

sur

face

1050

C11

0085

079

010

20?

996

15O

ccip

ital

dep

th: f

rom

upp

erm

ost p

oint

of o

ccip

ut to

low

er s

ide

cond

yle

680

C67

048

064

559

8

16P

osit

ion

of c

ondy

le: d

ista

nce

from

upp

er c

ondy

lar

edge

to p

oste

rior

edg

e of

mas

tica

tory

su

rfac

e of

upp

er m

olar

alo

ng e

dge

565

515

17M

inim

um d

epth

of t

he z

ygom

atic

5483

56.5

71

18L

engt

h of

zyg

omat

ic a

rch

300

C58

040

248

044

6

19A

nter

ior

pala

tal w

idth

: dis

tanc

e be

twee

n m

olar

alv

eoli

at a

nter

ior

edge

s~

9754

20P

oste

rior

pal

atal

wid

th: d

ista

nce

betw

een

mol

ar a

lveo

li at

pos

teri

or e

dges

~15

011

9

21S

kull

wid

th a

t sup

raor

bita

l pro

cess

es87

010

4082

073

068

088

0

22W

idth

of o

ccip

ital

con

dyle

s28

422

520

826

722

6

23D

iam

eter

s of

fora

men

mag

num

, hei

ght ×

wid

th10

0 ×

72

90 ×

82

59 ×

83

78 ×

83

24A

ngle

bet

wee

n cr

est b

orde

ring

ant

erio

rly

low

er p

art o

f tem

pora

l fos

sa a

nd s

kull

mid

line

3855

4543

25A

ngle

of d

iver

genc

e of

tusk

alv

eoli

1522

2315

26H

eigh

t (14

) to

leng

th (

1) r

atio

of s

kull

80.0

63.2

91.8

76.0

75.9

68.3

27D

epth

(15

) to

wid

th (

9) r

atio

of o

ccip

ut12

2.8

157.

614

0.8

117.

813

4.6

28R

atio

of s

kull

wid

th a

t sup

raor

bita

l pro

cess

es (

21)

to o

ccip

ital

wid

th (

9)96

.010

1.5

95.1

92.6

111.

891

.5

29R

atio

of m

inim

um fo

rehe

ad w

idth

(12

) to

gre

ates

t for

ehea

d le

ngth

(5)

61.4

66.3

67.5

53.4

53.4



Tabl

e 3.

Mea

sure

men

ts o

f th

e lo

wer

jaw

of t

he

step

pe e

leph

ant

Mam

mut

hus

trog

onth

erii

(P

ohli

g, 1

885)

Mea

sure

men

ts, m

mU

st’�

Tark

a,sp

ecim

en I

AE

,no

. 18

Pya

tiry

zhsk

,sp

ecim

en M

P P

GP

I P

2002

.114

9 (S

hpan

sky

et a

l.,

2008

a)

Azo

v 1,

,

spec

imen

AM

Z,

no. K

P�2

1081

(Bai

gush

eva

and

Gar

utt,

198

7)

Azo

v 2,

,

spec

imen

AM

Z,

no. K

P�2

8689

(B

aigu

shev

a an

d T

itov

, 201

0)

Nov

ogeo

rgie

vsk

, (Z

akre

vs’k

a,

1935

)

Len

gth

of h

oriz

onta

l ram

us: h

oriz

onta

lly a

t lev

el o

f mol

ar a

lveo

li (w

itho

ut m

en�

tal p

rotu

bera

nce)

472

480

600?

660

Dep

th o

f hor

izon

tal r

amus

alo

ng a

nter

ior

edge

of a

lveo

lus

240

235

201.

5

The

sam

e al

ong

post

erio

r ed

ge16

917

118

715

8

Gre

ates

t thi

ckne

ss o

f hor

izon

tal r

amus

194

180

174

176.

520

0

Dis

tanc

e be

twee

n ho

rizo

ntal

ram

i at a

nter

ior

inte

rnal

edg

e of

alv

eoli

8570

65

Gre

ates

t dis

tanc

e be

twee

n bu

ccal

surf

aces

of h

oriz

onta

l ram

i (at

pos

teri

or e

dge)

577

640

620*

620

Min

imum

dis

tanc

e be

twee

n m

olar

s, a

nter

iorl

y68

6710

1.7

90

The

sam

e, p

oste

rior

ly20

018

531

026

0

Gre

ates

t dis

tanc

e be

twee

n ho

rizo

ntal

ram

i (ex

tern

ally

)28

227

534

0

Sym

phys

eal d

epth

112

9311

3.5

130

Ant

erop

oste

rior

sym

phys

eal d

iam

eter

(in

clud

ing

men

tal p

rotu

bera

nce)

150

160

156.

2

Len

gth

of m

enta

l pro

tube

ranc

e73

141

72

Dep

th o

f asc

endi

ng r

amus

484

485

C45

047

0

Gre

ates

t dia

met

er o

f asc

endi

ng r

amus

357

C44

4

Len

gth

of m

enta

l par

t of l

ower

jaw

: fro

m a

nter

ior a

lveo

lar e

dge

to e

nd o

f men

tal

prot

uber

ance

300

380

Dis

tanc

e fr

om m

enta

l pro

tube

ranc

e to

pos

teri

or s

ide

of a

rtic

ular

hea

d78

078

088

0C

810

Dis

tanc

e fr

om a

nter

ior

alve

olar

edg

e to

pos

teri

or s

ide

of a

rtic

ular

hea

d51

551

054

0/60

0C

490

Dis

tanc

e be

twee

n ar

ticu

lar

head

s (i

nter

nally

)33

639

3C

345

The

sam

e ex

tern

ally

~52

057

063

0C

515

Len

gth

× w

idth

of a

rtic

ular

hea

d96

× 9

285

.6 ×

93.

585

× 1

15C

84 ×

C

97

Rat

io o

f dis

tanc

e be

twee

n ar

ticu

lar

head

s ed

ges

to d

ista

nce

betw

een

exte

rnal

w

alls

of h

oriz

onta

l ram

i (%

)~

90.1

89.1

101.

6>

83.1

Ang

le o

f con

verg

ence

of l

ower

jaw

ram

i (at

ext

erna

l wal

ls)

~70

°60

°

* S

ee t

he

text

.

312


SHPANSKY et al.

canal is wide, trapezoidal, with a widened ventral part.The ventrocaudal edge of the vertebral centrum has aweak depression, which is distinct in the southern ele�phant (Garutt, 1954).

Table 7 shows measurements of cervical vertebrae3–7. The third and fourth vertebrae are trapezoidal inoutline, with a wider ventral part and narrowed dorsalpart. In ventral view, the centrum of these vertebrae isslightly convex. The transverse foramina are oval,extending dorsoventrally. The transverse rib processes(pr. costotransversarius) spread aside and do notdescend beyond the ventral part of the vertebral cen�trum. The spinal canal is triangular, with widelyrounded corners.

The transverse processes are the largest in the sixthvertebra; they are directed ventrolaterally, consider�ably projecting beyond the ventral edge of the vertebralcentrum. The distal parts of the transverse processes ofthe sixth vertebra have large tubercles on the caudalside, which are hooked, entering the ventral incisuresof transverse processes of the succeeding seventh ver�

tebra. The sixth vertebra has small pathological out�growths on the ventral side of the transverse processes.The cranial side of both transverse processes hasforamina (about 4 cm in diameter) probably formed inconnection with lifetime resorption of spongy tissue ofthe vertebra. The fourth and fifth vertebrae have simi�lar incisures in the same place, although they are lessdeveloped (particularly in the fourth vertebra). Similarincisures have previously been recognized in the tho�racic vertebrae of the elephant from Pyatiryzhsk(Shpansky et al., 2008a).

The ventral part of the centrum of the seventh cer�vical vertebra has facets on the caudal side for attach�ment of the first pair of ribs. The ventral part of the ver�tebral centrum is strongly widened in the transverseplane and the dorsal part is narrowed. The ventrallength of the vertebral centrum is considerably greaterthan the dorsal length. The spinal canal of the seventhvertebra is also triangular, with concave walls. Thewidth of cervical vertebrae in the region of the trans�verse processes is identical in all vertebrae from the

Table 4. Measurements (mm) of molars M3/m3 of Mammuthus trogontherii (Pohlig, 1885) from localities of the WestSiberian Plain and Europe

Locality, collectionno. (reference)

Crown length

Crown width

Crown height

Numberof plates

Mean length

of plate

Frequencyof plates

per 10 cm

Enamelthickness

Ust’�Tarka, specimen IAE,no. 18, (M3/m3), dex; sin –

Pyatiryzhsk, specimen MP PGPI, no. P2002.1149, (m3), dex; sin (Shpansky et al., 2008a)

*215; 225 85.5; 91.2 – 12 + ?; 13 + ? 17.5; 18 6 1.8–[2.5]–3.1

Azov 1, specimen AMZ no. KP�21081, (M3/m3), dex; sin (Baigusheva and Garutt, 1987)

Azov 2, specimen AMZ no. KP�28689, (M3/m3), dex; sin (Baigusheva and Titov, 2010)

West Runton, specimen 1996.181, M3; 1992.36 m3, dex; sin (Lister and Stuart, 2010)

Chembakchino, specimenKhM�10398, (M3/M3), dex; sin (Kosintsev et al., 2004)

Asino, specimen TOKM, no. 10300/3, (M3), dex; sin(Shpansky, 2000)

*215 98; 102 _ c16 17; 16 7; 7.5 3.0

Chernyi Yar, specimen PIN, no. 4874, (M3/m3), dex; sin (Dubrovo, 1966)

–

Or’ya River, specimen OF�909, (m3), dex; sin (Garutt, 1972)

c190; c195

c84; c75 13 + ? 6.5–8 1.8–2.1

* Incompletely erupting teeth; therefore, measurement concerns accessible part.

*189; 210203; 194

�� 94; 97.5104; 108�� 10 ?+

8.5; 7.5 ?+�� 17.4; 18.9

21.3; 23.4�� 6.2; 5.6

4.5; 4.3�� 2.3–[2.7]–3.2

2.7–[3.2]–4.0��

354; 372418

�� 104; 10591

�� 191; 189168

�� 20; 2122

�� 1917�� 5.3

5.9�� 3.2–1.8

2.4–2.2��

380; 385330

�� 120.5; 120110

�� 210; 204135

�� 2221�� 16

18.74�� 5.5

5.25�� 2.94

3.53��

387; c400430

�� c118104�� 196

167�� 22

21�� 6.52

5.25�� 2.7

2.0–[2.3]–2.7��

278315�� 85.7; 80

80�� 178; 181

145.3�� 20

22�� 13.4; 12.9

16.7�� 7.25; 7.5

5.75�� –

1.9��

.2.58; 252�� 105; 106

98; 97�� 20

18�� 7; 6.5

6; 5.5�� 2.5

2.5��



second to seventh; with the caudal articular surfacegradually increasing in width and the vertebral cen�trum increasing in height.

The first, fifth, ninth, and 18th thoracic vertebraeare preserved. Vertebrae 9–11 are inaccessible formeasurements. The thoracic vertebrae are slightlysmaller than those of the skeleton from Pyatiryzhsk(Table 8). The angle of inclination of the neural spinesof thoracic vertebrae varies from 15° (first thoracic ver�tebra) to 45° (15th–18th). The neural spines increasein length from the first to fourth (550 mm) thoracicvertebra (in elephants from Pyatiryzhsk and Novo�

georgievsk, the neural spine of vertebra 5 is the longest(555 and 475 mm, respectively); in the southern ele�phant from Nogaisk, the longest spine is in vertebra 4(712 mm) (Zakrevs’ka, 1935; Garutt, 1954; Shpanskyet al., 2008a). Beginning from vertebra 5, the neuralspine decreases in length, becoming 180 mm long inthoracic vertebra 18. The spinal canal changes inshape from almost triangular to oval (from vertebra 5in caudal direction), with the width exceeding height.In the last thoracic vertebrae, it becomes almost circu�lar. The costal facets and transverse processes are dis�placed gradually dorsally from the ventral position and

Table 5. Measurements (mm) of the atlas of Mammuthus trogontherii (Pohlig, 1885)

MeasurementUst’�Tarka, specimen

IAE, no. 18

Pyatiryzhsk, specimen MP

PGPI, no. P2002.1149

(Shpanskyet al., 2008a)

Chembakchi�no, specimenKhM�10398 (Kosintsev

et al., 2004)

Azov 1, , specimen AMZ, no. KP�21081

(Baigusheva and Garutt, 1987)

Distance between edges of ala atlantis 427 475 C390 457

Greatest height of vertebra 253 230 208.5 268

Distance between external edges of fac. art. cranialis 278 270 243 269

Distance from edge of fac. art. cranialis to edge of ala atlantis 122 76

Minimum width of canalis vertebralis (in middle part) 60 67.5 64

Greatest width of canalis vertebralis (in upper part) 93 92 82 94

Height of canalis vertebralis 98 125 101

Distance between edges of fac. art. caudalis 237 245 199

Vertebral length at arcus dorsalis 107 95 68 117

Width of arcus dorsalis 172 195 173 167

Table 6. Measurements (mm) of the epistropheus of Mammuthus trogontherii (Pohlig, 1885)

MeasurementUst’�Tarka,

specimen IAE, no. 18

Pyatiryzhsk, specimen MP

PGPI, no. P2002.1149 (Shpansky et al.,

2008a)

Chembakchino, specimen

KhM�10398 (Kosintsev et al.,

2004)



West Runton, specimen

1996.181 (Lister and Stuart, 2010)

Greatest height 328 – С275 – 326

Greatest width at transverse processes 346 360 374

Width anterior to external edges of transverse foramina

274 – С208

Width of anterior articular surface 246 260 209.5 249 240

Height of posterior articular surface 164 170 138 – –

Width of posterior articular surface 183 200 162 – –

Height of spinal canal (anteriorly) 61 – 65.5 – –

Width of spinal canal 73 – 66.2 – –

314


SHPANSKY et al.

occupy a dorsolateral position on the vertebral centraafter vertebra 5. In the first two vertebrae, the ventrallength of the centrum is greater than the dorsal length;in succeeding vertebrae, the dorsal length of the verte�bral centra gradually becomes greater than the ventrallength and the greatest difference is observed in the lastthoracic vertebrae.

The neural spine of the last lumbar vertebra is atmost 150 mm long. The ventral side of the vertebralcentrum has pathological outgrowths in the shape ofspinate and hooked processes, osteophytes. The out�growths are directed both cranially and caudally,embracing ventrally the cranial part of the sacrum(Pl. 11, fig. 4). The development of osteophytes (bonespurs) is probably connected with the senile degenera�tive–dystrophic processes in bone tissue of the verte�bral centra; however, it is hardly probable that theycaused the death of the elephant, because they developover a rather long time. Similar osteophytes areobserved on the vertebrae of the cave bear from IrgikCave (Tasnadi�Kubacska, 1962).

The sacrum consists of five fused vertebrae. In thedorsal plane, they form a vaulted bend. On the cranialside, the sacrum is 290 mm wide at the wings; on theventral side, it is 420 mm long; the caudal width at thewings is 210 mm; the total cranial height of the sacrumis 160 mm; the centrum of the cranial sacral vertebra is110 mm wide; the cranial articular facet of the cranialsacral vertebra is 200 mm wide; the caudal articularfacet of the caudal vertebra is 100 mm wide. The sacrumof the female from Azov 2 (AMZ, no. KP�28689), alsoconsisting of five fused vertebrae, is somewhat larger;on the ventral side, it is 432 mm long, 310 mm widecranially at wings, and 175 mm wide caudally. In theelephant from West Runton, the sacrum includes sixfused vertebrae (due to fusion with the first caudal ver�tebra), so that it is 565 mm long on the ventral side and322 mm wide at the wings (Lister and Stuart, 2010),

which is the greatest value of this parameter. In thesteppe elephant from Novogeorgievsk, like the mam�moth from the Berezovka River, the number of fusedsacral vertebrae is reduced to four (Zalensky, 1903;Zakrevs’ka, 1935).

Seventeen left and 12 right ribs are preserved. Theanterior ribs are massive, with gradually expandingdistal ends. The medial surface is flattened and the lat�eral surface is slightly convex. The proximal end isturned medially relative to the sagittal plane. The firstrib is 700 mm long and its distal end is 185 mm wide.In the elephant from Ust’�Tarka, ribs curve veryweakly and smoothly. One of its longest ribs is thefourth, it is 1000 mm long along the arc chord and1200 mm long along the curvature (in the elephantfrom Novogeorgievsk, this parameter is 1210 mm inthe ninth rib: Zakrevs’ka, 1935); the rib head is 88 mmin longitudinal diameter.

Two segments of the sternum are preserved; theanterior segment is 360 mm long and 165 mm highanteriorly and 170 mm high posteriorly; its greatestwidth is 115 mm. The posterior segment is 288 mmlong and 147 mm high in the anterior part and 109 mmhigh posteriorly; the greatest width is 100 mm.

Pelvis (Table 9). The iliac wings are very wide andflat, the ventrolateral edges curve smoothly anteriorly(Pl. 11, fig. 1). The incisure of the acetabulum openson a flattened surface rather than directly in the ovalforamen. The distance from the incisure edge to theedge of the oval foramen is 55 mm. Dubrovo (1982)proposed that this position of the incisura acetabuli isa diagnostic character of M. primigenius; however, thesame incisure pattern of the acetabulum is character�istic of the elephant from Chembakchino (Kosintsevet al., 2004) and the female from Azov 2 (specimenAMZ, no. KP�28689). The measurements and shapeof the ilia and the indices proposed by Lister (1996b)suggest that this is a male skeleton. The indices of the

Table 7. Measurements (mm) of cervical vertebrae of Mammuthus trogontherii (Pohlig, 1885)

Measurement

Vertebra / locality

third fourth fifth sixth seventh

Ust’�Tarka

Ust’�Tarka Pyatiryzhsk Ust’�

TarkaUst’�Tarka Pyatiryzhsk Ust’�

Tarka Pyatiryzhsk

Vertebral height from ventral edgeof centrum to dorsal surface of prezygapophyses

252 278 C265 310 320 297 C350 305

Greatest width at transverse processes 370 370 C275 370 C365 262 370 380

Height�to�width ratio of vertebral centrum, anteriorly

185/180 200/180 195/195

Height�to�width ratio of vertebral centrum, posteriorly

/185 /186 195/193 /185 /195 200/200 /197 185/175

Height of spinal canal (cranially) C55 56 63

Width of spinal canal 95 117 132



Tabl

e 8.

Mea

sure

men

ts (

mm

) of

th

e th

orac

ic a

nd

lum

bar

vert

ebra

e of

Mam

mut

hus

trog

onth

erii

(P

ohli

g, 1

8)

No. of vertebra

Locality

Mea

sure

men

t

1. Widthat transverseprocesses(greatest)

2. Cranialwidthof vertebralcentrum

3. Cranial heightof vertebralcentrum

4. Caudal widthof vertebralcentrum

5. Caudal heightof vertebralcentrum

6. Widthof spinal canal(anteriorly)

7. Heightof spinalcanal (anteriorly)

8. Lengthof neuralspine alonganterior edge

9. Minimumwidthat articular costalfossae (anteriorly/posteriorly)

10. Lengthof vertebralcentrum dorsally/ventrally

IU

st’�

Tar

ka39

0C

84C

5938

0C

72/8

1P

yati

ryzh

skC

325

187

170

175

175

120?

115/

140

60/8

5II

Ust

’�T

arka

390

480

Pya

tiry

zhsk

410

170

162

160

163

9377

485

150/

150

60/9

0

Azo

v 2,

*

485

III

Ust

’�T

arka

360

520

Pya

tiry

zhsk

382

160

160

140

165

8863

?55

016

0/15

578

/82

Azo

v 2,

30

515

615

2.5

155

156

8553

440

120/

142

65/5

6

IVU

st’�

Tar

ka35

055

0P

yati

ryzh

sk36

014

515

514

116

275

5554

015

0/14

085

/78

Azo

v 2,

31

514

015

915

915

587

35,5

535

141/

127

63/6

3

VU

st’�

Tar

ka34

552

0P

yati

ryzh

sk36

514

816

015

016

583

4555

514

0/14

590

/83

Azo

v 2,

29

513

414

616

115

682

7849

010

7/10

269

/66

XII

Ust

’�T

arka

355

Azo

v 2,

*

C37

5

XII

IU

st’�

Tar

ka34

580

/80

XIV

Ust

’�T

arka

330

82/7

8

Azo

v 2,

*

224

107

126.

513

213

2.5

4965

365

/76

85/7

8

XV

Ust

’�T

arka

279

138

280?

85/7

9

Azo

v 2,

*

154

136

/159

XV

IU

st’�

Tar

ka26

814

326

090

/82

Azo

v 2,

13

113

8.5

150

138

7462

99.5

/92.

5

XV

IIU

st’�

Tar

ka25

813

024

590

/81

Pya

tiry

zhsk

*C

230

132

130

145

130

6058

310

120/

120

90/8

5

Azo

v 2,

13

514

214

114

3.5

6789

.5/9

1

XV

III

Ust

’�T

arka

235

139

180

104/

84

Azo

v 2,

12

913

513

214

750

6622

298

/84

VU

st’�

Tar

ka31

512

5~

150

81/

Azo

v 2,

29

916

312

9.5

182

125

106

3385

/92

*S

eria

l num

ber

of v

erte

bra

is d

eter

min

ed a

ppro

xim

atel

y.

316


SHPANSKY et al.

Table 9. Measurements of pelvic bones of Mammuthus trogontherii (Pohlig, 1885)

Measurement (right/left), mmUst’�Tarka, , specimen IAE,

no. 18

Pyatiryzhsk, , specimen

P2002.1149 (Shpansky et al.,

2008a)

Chembakchino, , specimen

KhM�10398(Kosintsev et al.,

2004)

Azov 2, ,specimen AMZ,no. KP�28689

1 Horizontal diameter of glenoid cavities

199 190 190

2 Vertical diameter of glenoidcavities

202 215 195 208

3 Greatest width at ilia (distance between wings)

1550 1700 1820

4 Width between external edgesof acetabulum

800 770

5 The same between internal edges 460 420 500

6 Greatest width of iliac wing 1000/900 840 1050/990

7 Symphyseal length 520 560

8 Length of ischiadic foramen 240 175? 202 237/254

9 Width of ischiadic foramen 110 120? 107 133/127

10 Distance from upper edge of ilium to middle part of glenoid cavity

760 380? 860/790

11 Distance between external edges of ischiadic bones

440 470 600

elephant from Ust’�Tarka differ from that of femalesfrom Azov 2 and Edersleben (Fig. 4, Table 10). Thepelvis of the elephant from Ust’�Tarka differs addi�tionally from that of females in the flattened wings ofthe ilia and the straight crest of the iliac wing, as inM. primigenius; the pelvis of the male from Azov is notpreserved; in the female pelvis from Azov, “… the crestof the iliac wing is convex. The wing surface is rough,concave on both sides” (Baigusheva and Titov, 2010,p. 20). The greater ischiadic foramen (apertura pelvis)is circular; its width is less than the diagonal heightmeasured from the symphysis to the lowermost pointof connection with the sacrum. Such a ratio is typicalfor females, whereas males usually show an inverseratio. The width of the greater ischiadic foramen couldhave been limited by the discontinuance of growth ofthe pubis and ischium at the point of maturity, whereasthe foramen height could have increased due to thegrowth of the ilia (Tikhonov, 1996).

The femora have well adherent epiphyses; the epi�physeal sutures are hardly discernible. The diaphysealwidth is considerably greater than the anteroposterior

diameter (Table 11). The neck of the head is relativelyshort and massive, directed medially at an angle of 50°.At the supraglenoid tubercles, the distal end is muchwider than the distance between the external edges ofthe medial and lateral condyles. The articular surfacefor the patella is wide and slightly concave, somewhathigher on the medial side than laterally.

The tibia are almost straight; in the dorsal plane,the proximal part is trapezoidal, so that the plantarside is wider. The plantar side is concave in the upperone�third, limited laterally by sharp crests (lineaemuscularum). The roughness of the tibia is triangular,with a pointed distal end. The main part of the diaphy�sis is triangular in cross section, with a well�pro�nounced crest (crista tibiae). The lateral and medialwalls are flat; in the proximal part, the medial wall hasa small rough depression. The tibiae of the elephantfrom Ust’�Tarka are relatively short and massive, con�siderably inferior in absolute length to the bones ofmales from Azov 1 and Odessa, but superior in the dia�physeal width (Table 11). The lateral articular surfaceof the proximal epiphysis is positioned much lower

E x p l a n a t i o n o f P l a t e 1 1

Figs. 1–4. Mammuthus trogontherii (Pohlig 1885), specimen IAE, no. 18; Novosibirsk Region, Ust’�Tarka; Middle Neopleis�tocene, Tobolsk horizon: (1) pelvis and sacrum, anterior view; (2) left femur with mounted patella, anterior view; (3) distal regionof left hind leg, including mounted tibia, fibula, and foot, anterior view; (4) posteriormost lumbar vertebra, with well�developedosteophytes (indicated by arrows), ventral view.



Plate 11

10 cm

5 cm

10 cm 10 cm

1

23

4

318


SHPANSKY et al.

than the medial facet. The facet for the proximal endof the fibula is relatively small, oval.

In the left fibula, the diaphysis is only partially pre�served; and in the right counterpart, it is broken. Thebones are slightly bent. On the medial side, the distalpart of the diaphysis is strongly flattened and expandedin the sagittal plane. The reconstructed right bone is731 mm long. Its proximal epiphysis is 81 mm wideand 56 mm in sagittal diameter; the diaphysis is47.5 mm wide in the middle; the distal epiphysis is 78mm wide and 138 mm in sagittal diameter.

The patella is oval; its cranial surface is gently con�vex, rough, covered with many small nutrient foram�ina. The patella of the elephant from Ust’�Tarka is rel�atively small; it is larger than in the elephant fromChembakchino but considerably smaller than that oflarge males from Pyatiryzhsk and Azov 1 (Table 12).

SKELETON PROPORTIONS

The skeleton of M. trogontherii from Ust’�Tarkamounted on a metal frame shows the following mea�surements. The length from the anterior surface of thepremaxillae to the posterior edge of the ischium is390 cm. At both the most projecting skull part and the

apex of the neural spine of the third thoracic vertebra(at the withers, the highest dorsal point), the skeletonis approximately 350 cm high (Fig. 2). The height ofthe mounted skeleton at the withers is estimated basedon the forelimbs reconstructed using the skeleton ofthe elephant from Novopetrovsk (specimen PIN,no. 4464; Moscow Region, Istrinskii District), whichis similar in size. At the dorsal edge of the pelvis, themounted skeleton of the elephant from Ust’�Tarka is293 cm high. The sum of lengths of the foot, tibia,femur, and ilium (from the acetabulum to dorsaledge), is 323 cm. The proportions of the hind limbwere calculated based on this parameter, since it allowscomparisons with other skeletons, including those notmounted. The difference in the above dimensions fol�lows from the losses in the pelvic and knee articula�tions and mounting the skeleton in a moving posture(Fig. 2). The hind limb proportions in the elephantfrom Ust’�Tarka are close to the mean values of steppeelephants. At the sacrum, males are 293–374 cm highand females are 300–342 cm high. The femur length is40.3% and the tibia length is 24.1%. Small differencesare caused by the underestimated distance from thefemoral head to the dorsal edge of the ilia (75 cm) andthe large foot (the total height of mounted foot bones

Table 10. Characters of sexual dimorphism in pelvic bones of elephants (after Lister, 1996, modified and supplemented):measurements: (1) greatest width of the pelvis at wings; (2) diagonal height of the greater ischiadic foramen from the sym�physis to lower point of connection with the sacrum; (3) maximum horizontal width of the greater ischiadic foramen; (4)width iliac wing from pelvic tubercle to the nearest point of the greater ischiadic foramen; (5) minimum width of iliac neckparallel to anterior plane of the neck

Species, sex, and localityMeasurement, mm Index

1 2 3 4 5 5/1 3/1 3/2 3/4 3/5 2/5

Ahidiskodon meridionalis,

, Valdarno, n = 3

1600–1880

400–470

440–540

– 251–284

0.141–0.169

0.275–0.287

1.04–1.15

– 1.63–1.95

1.48–1.87

A. meridionalis, , Valdarno, n = 2

1300–1380

450 440–470

– 156–170

0.113–0.131

0.319–0.362

0.98–1.04

– 2.76–2.82

2.65–2.88

A. meridionalis, ,Rodionovo*

1620 – 452 610 – – 0.279 – 0.741 – –

Mammuthus trogontherii, , West Runton

C1760 – 490 – 244 0.139 0.278 – – 2.01 –

Mammuthus trogontherii, , Ust’�Tarka

1550 520 455 630 253 0.163 0.294 0.88 0.720 1.80 2.06

M. trogontherii, , Azov 2 1820 723 583 610 220; 209

0.121; 0.115

0.320 0.806 0.956 2.65; 2.78

3.28; 3.45

M. trogontherii, ,Edersleben

1540 630 570 630 192 0.125 0.370 0.90 0.904 2.97 3.28

M. primigenius, , n = 11 1120–1600

330–480

410–510

460–620

182–208

0.133–0.186

0.306–0.384

1.0–1.24

0.788–1.0

1.92–2.64

1.69–2.42

M. primigenius, , n = 2 1170–1340

390 440–540

400–452

154–194

0.132–0.145

0.377–0.403

1.13 0.973–1.35

2.78–2.86

2.53

* Initial measurements of pelvic bones of A. meridionalis, specimen BX�2120 from Rodionovo are given after Maschenko et al. (2011).



Tabl

e 11

.M

easu

rem

ents

of l

ong

hin

d li

mb

bon

es o

f Mam

mut

hus

trog

onth

erii

(P

ohli

g, 1

885)

from

Wes

tern

Sib

eria

an

d E

aste

rn E

urop

e

Mea

sure

men

ts (

mm

),in

dice

s (%

)F

emur

Tib

ia

Loc

alit

y

Ust’�Tarka, IAE, no. 18

Chembakchino,KhM�10398(Kosintsev et al., 2004)

Pyatiryzhsk, P2002.1149(Shpansky et al., 2008a)

Odessa (Yatsko, 1948)

Azov 1, AMZ, no.KP�21081 (Baigushevaand Garutt, 1987)

Azov 2, AMZ, no.KP�28689 (Baigushevaand Titov, 2010)

West Runton, no. 1996.181.22(Lister and Stuart, 2010)

Ust’�Tarka, IAE, no. 18

Chembakchino,KhM�10398(Kosintsev et al., 2004)

Azov 1, AMZ, no.KP�21081 (Baigushevaand Garutt, 1987)

Azov 2, AMZ, no.KP�28689 (Baigushevaand Titov, 2010)

Odessa (Yatsko, 1948)

West Runton, no.1996.181.27(Lister and Stuart, 2010)

1. L

engt

h13

0212

3015

0014

80–

1380

1425

777

645

890

825

920

830

2. D

iaph

ysea

l wid

th18

011

117

2?20

2C

230

173

180

132

110.

512

712

012

513

2

3. D

iaph

ysea

l dia

met

er11

594

–11

2–

–11

988

123

107

110

–

4. G

reat

er h

ead

diam

eter

191

–21

018

823

019

521

228

323

628

526

628

229

9

5. T

ran

sver

se h

ead

diam

eter

––

––

––

246

187

217

220

245

6. W

idth

of d

ista

lep

iph

ysis

280

243

290

326

340

291

323

206

196

255

221

221

248

7. T

ran

sver

se d

iam

eter

of d

ista

l epi

phys

is30

6–

320

–27

930

430

816

215

817

217

315

118

3

Inde

x of

dia

phys

is(2

: 1)

13.8

9.0

13.6

12.5

12.6

17.0

17.1

14.3

14.5

13.6

15.9

Pro

xim

al in

dex

(4 :

1)–

––

––

––

36.4

36.6

32.0

32.2

30.7

36.0

Dis

tal i

nde

x (6

: 1)

21.5

19.8

19.3

22.0

–21

.122

.726

.530

.428

.726

.824

.029

.9

320


SHPANSKY et al.

is 40 cm). In the elephant from Ust’�Tarka, forelimbbones are not preserved; however, based on the ratio ofthe skeleton height at the withers to the height in thesacral region, which was previously obtained by us(Shpansky et al., 2008b), it is possible to estimate theskeleton height at the withers. In the steppe elephant,this ratio is 1.1 and, hence, as the height at the upper�most point of the pelvis is 323 cm, the height at thewithers of the skeleton from Ust’�Tarka is estimated as355 cm. These measurements are relatively small for

males; thus, the elephant from Ust’�Tarka is consider�ably inferior in size to the giants from Pyatiryzhsk(424 cm high at the withers), Azov 1 (420 cm), WestRunton (400 cm), and Gelaineo (389 cm) and supe�rior to females from Novogeorgievsk (339 cm) andEdersleben (Table 13). During the animal’s life, theelephant from Ust’�Tarka could have been about370 cm high at the withers, including soft tissues.

The ribs are only slightly greater in absolute sizethan ribs of M. primigenius. Free forelimb was about

Table 12. Measurements of patella of Mammuthus trogontherii (Pohlig, 1885)

Measurement, mmUst’�Tarka,

specimen IAE, no. 18

Pyatiryzhsk, specimen

P2002.1149 (Shpansky et al.,

2008a)

Chembakchino, specimen

KhM�10398 (Kosintsev et al.,

2004)

Azov 1,specimenKP�21081


West Runton, specimen 1995.105

(Lister andStuart, 2010)

1. Greatest length 148.5 179 143 171 158

2. Greatest width 139 154 112 150 127

3. Greatest diameter 102 112 84 108 –

Table 13. Estimates of skeleton height of steppe elephants based on cylindrical limb bones (mm)

Locality, sex

Humerus(30.4% of skeleton height at withers)

Ulna(23.2% of skeleton

height at the withers)

Femur(40.5% of skeleton height at sacrum)

Tibia(24.4% of skeletonheight at sacrum)

bone length

estimated skeleton height

bone length


bone length


bone length


Novogeorgievsk, 980 3224 850 3664 1210 2988 690 2828

Gelsenkirchen, 1190 3914 950 4095 1300 3210 760 3115

Edersleben, 1066 3507 802 3457 C1277 3153 685 2807

*West Runton, 1240 4079 920 3965 1425 3518 830 3402

Odessa, 1230 4046 980 4224 1480 3654 920 3770


1290 4243 c950 4095 1480? 3654 900 3689


1160 3816 880 3793 1380 3407 825 3381

Chembakchino, , specimen KhM�10398

1010 3322 C570 1230 3037 645 2643

Pyatiryzhsk, ,specimen P2002.1149

1320 4342 940 4052 1500 3704 – –

Ust'�Tarka, ,specimen IAE, no. 18

– – – – 1302 3215 777 3184

* Lister and Stuart (2010) estimated the height at shoulders of the elephant from West Runton with soft tissues as 390 cm.



160 cm long (in the male from Azov 1, it is about180 cm long).

TAXONOMIC POSITION

The taxonomy of mammoth�like elephants is basedon dental morphology of the last tooth generation(M3). Additional data for the diagnosis are providedby the morphology of the skull, including lower jaw(Garutt, 1954, 1998; Lister, 1996a). Based on theteeth, it is impossible to determine with certainty thetaxonomic position of the elephant from Ust’�Tarka,because they are heavily worn. The controversial ques�tion of the assignment of all Euroasian mammoth�likespecies to the genus Mammuthus (according to theWest European colleagues) or division into two genera(as is usually accepted by Russian researchers) isbeyond the scope of the present study.

The skull and lower jaw morphology of the ele�phant from Ust’�Tarka is transitional between the gen�era Archidiskodon and Mammuthus, as is typical forM. trogontherii (Fig. 5). The general questions of theevolutionary trend in changes of the skull and lowerjaw of A. meridionalis, M. trogontherii, and M. primige�nius have been investigated in detail (Maglio, 1973;Lister, 1996a). The skull and lower jaw of the specimenconsidered here (Figs. 5–7) are much closer in mor�phology to the woolly mammoth: (1) the skull is highand shortened in the sagittal plane, with a taperingapex; (2) the plane of occipital bones is convex; (3) theoccipital condyles relatively poorly project beyond theoccipital plane; (4) the forehead plane is almost even,its relative width (the ratio of the minimum foreheadwidth to occipital width) is 51.5%, which is character�istic of the genus Mammuthus, while in Archidiskodon,the relative forehead width is 29–47.1% (Garutt,1998); (5) the forehead width at the supraorbital pro�cesses is greater than the occipital width; (6) the loweredge of the nares is at the level of the orbital midline;

(7) the distance between the lateral edges of articularprocesses of the lower jaw is less than the distancebetween the lateral walls in the region of jaw angles;(8) the horizontal ramus of the lower jaw is deep, thesymphyseal region is short; the ascending ramus iswidened in the region of the coronoid process; thecrest of the coronoid process is well developed andraised.

The morphology of the premaxilla approaches thatof M. primigenius; therefore, at the alveolar level, thetusks are directed downwards and, distally, curve ante�riorly. In the southern elephant, they are directed ven�trally and anteriorly just at the alveolar level. The pre�maxillae of the elephant from Ust’�Tarka are relativelylong; the length�to�width ratio at the level of theinfraorbital foramina is 171.7%. In Archidiskodon, thisratio is 147.3–176.9% and, in Mammuthus, it is163.5–209.7% (Garutt, 1998). The region near thenares (the proximal end of the premaxillae, particu�larly the nasal process) and supraorbital processes ofA. meridionalis strongly project beyond the foreheadplane and beyond the nasal surface of the premaxilla,making inflated the middle skull part in projection onthe sagittal plane (Figs. 5a, 6a). In M. trogontherii andM. primigenius, the region near the nares and supraor�bital processes is in the forehead plane; at the sametime, M. trogontherii retains a weak longitudinal con�cavity on the frontal surface. The nares of A. meridion�alis are narrowed crescentic, 443–620 mm wide, 71–120 mm high, and the height index is 13.9–21.9%. InM. trogontherii, it is relatively high (Table 2), crescen�tic, but its lateral margins are extended downwards(Pl. 10, fig. 1); the height index is 25.1–28.6%. Thenares reach the maximum height in M. trogontheriichosaricus (155–195 mm) at 460–532 mm of width;the height index is 29.1–42.4%. In M. primigenius, thenares are high, widely oval, with round lateral edges; itis 325–472 mm wide, 80–150 mm high; the heightindex is 24.2–34.5%. The characteristics listed may

(a) (b) (c)

P�F

Fig. 5. Comparison of skulls of elephants of the mammoth�like lineage (lateral view): (a) Archidiskodon meridionalis, specimenIGF, no. 1054; Italy, Upper Valdarno (Garutt, 1954; Palombo and Feretti, 2005); specimen SM, no. 14120, lower jaw; Georgievsk(Garutt and Safronov, 1965); (b) Mammuthus trogontherii, specimen IAE, no. 18; Novosibirsk Region, Ust’�Tarka; specimenPIN, no. 4874, lower jaw; Volgograd Region, Chernyi Yar (Garutt, 1954); (c) M. primigenius, neotype ZIN, no. 2710; TaimyrPeninsula (Garutt and Tikhonov, 2001). Arrows show directions of morphological changes in the skull and lower jaw of mam�moth�like elephants in phylogeny.

322


SHPANSKY et al.

(a) (b) (c)

50 cm

Fig. 6. Skulls of elephants of the mammoth�like lineage, lateral view: (a) Archidiskodon meridionalis, Italy, Sauvignon (Regianiand Sala, 1994); (b) Mammuthus trogontherii, specimen IAE, no. 18; Novosibirsk Region, Ust’�Tarka; (c) M. primigenius, speci�men PM TGU, no. 54; Tomsk Region.

result from adaptation to arud climate, with the maxi�mum indices of the nasal opening height inthrogonthere elephant, which dwelt during the maxi�mum claxiation and in M. primigenius characteristic ofthe late Late Neopleistocene, with the maximumaridity in the latter half.

A characteristic feature is changes in the lower jaw,i.e., the shortened mental protuberance of the sym�physis, accompanied by an increase in the relativedepth of the horizontal ramus and expansion of thearea of the coronoid process (Figs. 5b, 7b). A signifi�cant expansion of the plane of the ascending ramusand rostral displacement of the anterior angle of thecoronoid process are similar to that of M. primigenius.A distinctive feature of the steppe elephant is theoblique angle of the lower jaw in projection on the sag�ittal plane, which results in a zigzag line and strong

caudal bulge of the upper one�third of the ascendingramus (Fig. 7b).

Table 14 shows the parameters of preserved distalparts of M3/m3.

A straight crest on the iliac wing, which was previ�ously regarded as a diagnostic character of M. primige�nius (Garutt, 1954; Dubrovo, 1982) is observed in theskeleton of the steppe elephant from Ust’�Tarka and inthe elephant Archidiskodon meridionalis cf. gromovifrom northeastern Italy (Reggiani and Sala, 1994). Inthe southern elephant A. meridionalis from Rodion�ovo, the crest of the iliac wing is gently bent(Maschenko et al., 2011). It should be noted that, inmammoth cubs below three–four years of age, thecrest of the iliac wing has a distinct curvature, which isparticularly well pronounced in the cubs younger thanone year of age (Shpansky and Pecherskaya, 2007).

Table 14. Comparison of teeth in Mammuthus trogontherii (Pohlig, 1885) and Archidiskodon meridionalis (Nesti, 1825)

Plate frequency per 10 cm Enamel thickness, mm

M. trogontherii A. meridionalis M. trogontherii A. meridionalis

Ust’�Tarka, specimen IAE,

no. 18

Extreme valuesin other skeletons

(see Table 4)

Shpansky, 1999; Maschenko et al.,

2011

Ust’�Tarka, specimen IAE,

no. 18

Extreme valuesin other skeletons

(see Table 4)

Shpansky, 1999; Maschenko et al.,

2011

M3 5.6–6.2 5.3–6.52 4.5–5.5 2.3–(2.7)–3.2 1.8–(2.7)–3.2 3.0–3.8

m3 4.3–4.5 5.25–6 4.5–5 2.7–(3.2)–4.0 2.0–(2.5)–3.53 2.5–4.0



Subsequently, the wing crest is straightened. Thus,according to our data, a straightened crest on the iliummay not be unequivocally taken for a distinctive char�acter of M. primigenius, although a rounded crest ofthe ilium has not been recorded in adult woolly mam�moths. In our opinion, the presence of such a structureof pelvic bones in the specimen from Ust’�Tarkamakes it closer to later members of the mammoth�likelineage.

Thus, the skull and lower jaw shape and features ofpelvic bones suggest the assignment of specimen IAE,no. 18 to M. trogontherii.

CONCLUSIONS

Skeletons of M. trogontherii and their fragments arerather rare in Western Siberia. All of them are confinedto the alluvial beds of the Irtysh River (Chembakchin�skii Yar, Pyatiryzhsk) and its tributary, the Om River(Ust’�Tarka). The skeleton from Ust’�Tarka is one ofthe smallest in absolute measurements and compara�ble to skeletons from Novogeorgievsk and Gelsen�kirchen (3.34–3.5 m high at the withers) (Shpanskyet al., 2008b). It is smaller than the female skeletonfrom the Kagal’nitskii quarry (Azov 2) (Baigushevaand Titov, 2010). Certain anatomical features of bones(such as obliteration of epiphyseal sutures, large tusks,

mental protuberance of the lower jaw, and morphologyof pelvic bones) suggest that the skeleton from Ust’�Tarka belongs to a male about 55–60 years of age.Functioning teeth in its jaws are heavily worn molarsof the last generation (M3/m3).

Presently known West Siberian skeletons ofM. trogontherii are of approximately the same geologi�cal age and connected with deposits of the TobolskHorizon dated the beginning of the Middle Neopleis�tocene. The date of 550–600 ka reported for the bedsenclosing the elephant skeleton from Chembakchino(Kosintsev et al., 2004) is doubtful because of mor�phological features and preservation of availableremains. The skeletons from Europe are somewhatmore ancient, come from the Early Neopleistocenebeds. The steppe elephant is an intrazonal species andits remains (usually teeth) found in Western Siberiaalso occur in the Early Neopleistocene beds in thecomposition of the Vyatka Faunal Assemblage. In theIrtysh Region near Pavlodar (and in Kazakhstan as awhole), the steppe elephant is recorded in the Kosh�kurgan (Early Neopleistocene) and Irtysh (terminalEarly Neopleistocene–basal Middle Neopleistocene:Lebyazh’e and Zhanaul formations of the IrtyshRegion near Pavlodar) faunal assemblages (Shpanskyet al., 2007; Shpansky, 2009).

(a) (b) (c)

20 cm

(d) (e) (f)

Fig. 7. Lower jaws of mammoth�like elephants, buccal view: (a, d) Archidiskodon meridionalis: (a) specimen SM, no. 14120;Georgievsk (Garutt and Safronov, 1965); (d) Italy, Oriolo (Palombo and Ferretti, 2005); (b, e) Mammuthus trogontherii: (b) spec�imen IAE, no. 18; Ust’�Tarka; (e) specimen MP PGPI, no. 2002.1149�1; Pyatiryzhsk (Shpansky et al., 2008); (c, f) M. primige�nius, specimen PM TGU, nos. 1/17 (c), 1/56 (f); Tomsk Region.

324


SHPANSKY et al.

ACKNOWLEDGMENTS

We are grateful to the anonymous reviewer forperusal of the manuscript and help in structurizationof the data provided in the paper.

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